Foster and Wedel 2014 fig 3 - dorsals

Fig. 3. MWC 8028, Haplocanthosaurus dorsal vertebrae. A. Lateral view of dorsal centrum with bottom edge of lateral pneumatic fossa preserved. B. Dorsal view of same centrum as in A, showing the median septum between the paired lateral fossae. C. Lateral view of dorsal centrum with smaller segment of the lateral pneumatic fossa margin preserved. D. Dorsal view of same centrum as in C, again showing the median septum and paired lateral fossae. E. Lateral view of dorsal centrum with partial pleurocoel preserved. F. Cross-sectional (posterior) view of same dorsal as in E. G. Dorsal neural spines in lateral (top) and anterior or posterior (center, bottom) views. Scale bars = 10 cm.

Right on the heels of Aquilops last week, my paper with John Foster on the new specimen of Haplocanthosaurus from Snowmass, Colorado, was just published in Volumina Jurassica. I’ll have more to say about it later, but right now I am up against a deadline on a big project and I need to go work on that. I’m only popping up here to note two quick things.

First, if you’re not familiar with Volumina Jurassica – and I wasn’t, before this project – it’s a free-to-access* journal that publishes papers on all aspects of the Jurassic. The current issue is specifically dedicated to the Jurassic formations of the American West. There’s a lot of interesting stuff in there, but of special interest to SV-POW! readers will be the paper by Cary Woodruff and John Foster on the legendary and possibly apocryphal Amphicoelias fragillimus.

* But not truly open access since the journal claims to retain exclusive rights to distribute the papers. That seems like a quaint affectation now that the internet is here, but whatever – at least they let anyone download the PDF for free, which is primarily what I care about.

Foster and Wedel 2014 fig 4 - sacrum

Fig. 4. Sacra of Haplocanthosaurus.  A. MWC 8028, sacrum in right lateral view. B. MWC 8028, close-up of S4 and S5 centra highlighting pneumatic fossae. C. MWC 8028 with divisions between the vertebrae overlaid. D. CM 879, sacrum in right lateral view with divisions between the vertebrae overlaid. E. CM 572 in right lateral view, after Hatcher (1903c: plate 4). B–E are not shown at the same scale, scale bar for A = 20 cm. Note that the neural arches in CM 572 were restored during preparation, and the sacral neural spines as shown here are probably lower than they would have been in life.

Second, the figure resolution in the PDF of the Haplocanthosaurus paper is not stellar, so as is the case with almost all of our papers, the full-color, high-resolution figures are available at the paper’s page on the sidebar.

Gotta run.

For our previous posts on Haplocanthosaurus, go here; for those on Amphicoelias, including Mike’s very popular, “How big was Amphicoelias fragillimus? I mean, really?”, go here.

References

A while back, Ben Miller reminded me that when I posted about the old Yale “Brontosaurus” skull, I promised:

So how did the YPM come to make such a monstrosity? What was it based on? Tune in next time for the surprising details!

I told him at the time that I’d soon get around to writing a post. But before I did, he wrote a post on this himself: Bully for Camarasaurus. And it’s excellent. Go and read it!

I don’t have a lot to add to what Ben has written, except regarding this:

What Marsh had instead [when restoring the skull for his 1891 “Brontosaurus” reconstruction] were a few fragmentary bits of Camarasaurus cranial material, plus a snout and jaw (USNM 5730) now considered to be Brachiosaurus.

Here’s what Marsh came up with:

Marsh1891-plateXVI-Apatosaurus-skull-UNREVERSED

But what of the supposed Brachiosaurus skull that he used as a reference? It was finally described 107 years later by Carpenter and Tidwell (1998), in a paper that helpfully also lays out the history behind it. Here’s how it looks:

CarpenterTidwell1998-fig1

The skull was found by a crew under the supervision of M. P. Felch in the western part of his Quarry 1, Garden Park, Colorado. Felch reported it to O. C. Marsh in a letter of 8 September 1883. It was found by a meter-long cervical vertebra that probably belonged to Brachiosaurus “which was destroyed during attempts to collect it” (McIntosh and Berman 1975:196). [Of course, Felch and Marsh could hardly have been expected to identify this vertebra correctly, as Brachiosaurus would not be discovered and named for another twenty years (Riggs 1903), and the nature of its neck would not become apparent until Janensch (1914) described the related brachiosaurid Giraffatitan (= “Brachiosaurus“) brancai.]

The Felch skull, along with other material from the quarry, was shipped to Marsh at Yale in October of that year, and was initially assigned the specimen number YPM 1986. At that time it was only partially prepared, hence the rather poor resemblance between the restored version above and Marsh’s hypothetical “Brontosaurus” [= Apatosaurus] skull that was based on it.

It’s notable that Holland (1915) was quite certain that this was not a skull of Brontosaurus, and that a Diplodocus-like skull found with the A. louisae holotype belonged to it. It’s worth reading the skull section of his paper to see just how solid his reasoning was. And it’s extraordinary to think that Osborn’s power, all the way over in New York, was so great that he was able to successfully bully Holland, 370 miles away in Pittsburgh, into not putting the evidently correct skull on the Carnegie Museum’s Apatosaurus mount. That mount remained sadly headless until after Holland’s death.

Aaanyway, YPM 1986 was pretty much ignored after Marsh’s abuse of it as a reference for the Brontosaurus reconstruction’s skull. After Marsh’s death in 1899, much of the material collected by Felch was transferred to the Smithsonian (US National Museum of Natural History). The skull was among these specimens, and so was re-catalogued as USNM 5730.

As so often, it was Jack McIntosh who rediscovered this skull and recognised its true affinities. Some time after his tentative identification of the skull as pertaining to Brachiosaurus (presumably on the basis of its resemblance to that of Giraffatitan), Carpenter borrowed the skull, had it more fully prepared, wrote the description, and had a restored model constructed from casts of the preserved elements and models of the missing ones.

Carpenter and Tidwell (1998:fig. 2) also handily showed the restored Felch quarry skull alongside those of other sauropods:

CarpenterTidwell1998-fig2

By re-ordering the top row, we can see what a neat intermediate it is between the skulls of Camarasaurus (left) and Giraffatitan (= “Brachiosaurus” of their usage):

CarpenterTidwell1998-fig2-top-row-reordered

I provisionally accepted USNM 5730 as belonging to Brachiosaurus in my re-evaluation of 2009, and included it in my reconstruction (Taylor 2009:fig. 7):

Taylor (2007: figure 7). Skeletal reconstruction of Brachiosaurus altithorax. White bones represent the elements of the holotype FMNH P 25107. Light grey bones represent material referred to B. altithorax: the Felch Quarry skull USNM 5730, the cervical vertebrae BYU 12866 (C?5) and BYU 12867 (C?10), the "Ultrasauros" scapulocoracoid BYU 9462, the Potter Creek left humerus USNM 21903, left radius and right metacarpal III BYU 4744, and the left metacarpal II OMNH 01138. Dark grey bones modified from Paul's (1988) reconstruction of Giraffatitan brancai. Scale bar equals 2 m.

Taylor (2007: figure 7). Skeletal reconstruction of Brachiosaurus altithorax. White bones represent the elements of the holotype FMNH P 25107. Light grey bones represent material referred to B. altithorax: the Felch Quarry skull USNM 5730, the cervical vertebrae BYU 12866 (C?5) and BYU 12867 (C?10), the “Ultrasauros” scapulocoracoid BYU 9462, the Potter Creek left humerus USNM 21903, left radius and right metacarpal III BYU 4744, and the left metacarpal II OMNH 01138. Dark grey bones modified from Paul’s (1988) reconstruction of Giraffatitan brancai. Scale bar equals 2 m.

But as noted by Carpenter and Tidwell (1998:82), the lack of comparable parts between the Felch skull and the Brachiosaurus holotype (which remains the only definitive Brachiosaurus material) means that the assignment has to remain tentative.

What we really need is a more complete Brachiosaurus specimen: one with both a skull and good postcervical elements that let us refer it definitively to Brachiosaurus altithorax by comparison with the holotype. And heck, while we’re at it, let’s have a specimen with a good neck, too!

The real question remains: how did Marsh, using a brachiosaur skull as his basis, come up with this?

Marsh1891-plateXVI-Apatosaurus-skull-UNREVERSED

 

And stranger still, how someone at the Yale Peabody Museum — we don’t know who — used it, or more likely Marsh’s reconstruction, as a basis for this sculpture:

IMG_0517

 

The Yale mount didn’t go up until 1931 — the last of the Big Four Apatosaurus mounts after the AMNH, Carnegie and Field Museum, which is surprising as it was the first of those specimens to be found. So by the time the skull was sculpted, sauropod skulls were actually reasonably well known. It’s not clear quite how anyone working from a decent reconstruction of, say, a Camarasaurus skull — the one in Osborn and Mook (1921:figure 30), say — could come up with this monster.

The last thing to say is this: it does credit to the YPM that they display this historically important sculpture rather than hiding it away and pretending it never happened. For me, part of the fascination of palaeontology is seeing not just how organisms evolved through prehistory but how ideas evolved through history. It’s great that we can still see important mistakes, alongside their corrections (i.e. the new and lovely skull on the YPM Apatosaurus mount.)

 

References

  • Carpenter, Kenneth, and Virginia Tidwell. 1998. Preliminary description of a Brachiosaurus skull from Felch Quarry 1, Garden Park, Colorado. Modern Geology 23:69-84.
  • Holland, William J. 1915. Heads and tails: a few notes relating to the structure of the sauropod dinosaurs. Annals of the Carnegie Museum 9:273-278.
  • Janensch, Werner. 1914. Ubersicht uber der Wirbeltierfauna der Tendaguru-Schichten nebst einer kurzen Charakterisierung der neu aufgefuhrten Arten von Sauropoden. Archiv fur Biontologie, Berlin III, 1(1):81-110.
  • Marsh, O. C. 1891. Restoration of Triceratops (with plates XV and XVI). American Journal of Science, 3rd series 41(244):339-342.
  • McIntosh, John S., and David, S. Berman. 1975. Description of the palate and lower jaw of the sauropod dinosaur Diplodocus (Reptilia: Saurischia) with remarks on the nature of the skull of Apatosaurus. Journal of Paleontology 49(1):187-199.
  • Osborn, Henry Fairfield, and Charles C. Mook. 1921. Camarasaurus, Amphicoelias and other sauropods of Cope. Memoirs of the American Museum of Natural History, n.s. 3:247-387, and plates LX-LXXXV.
  • Riggs, Elmer S. 1903. Brachiosaurus altithorax, the largest known dinosaur. American Journal of Science 15(4):299-306.
  • Taylor, Michael P. 2009. A re-evaluation of Brachiosaurus altithorax Riggs 1903 (Dinosauria, Sauropoda) and its generic separation from Giraffatitan brancai (Janensch 1914). Journal of Vertebrate Paleontology 29(3):787-806.

 

Last night, I submitted a paper for publication — for the first time since April 2013. I’d almost forgotten what it felt like. But, because we’re living in the Shiny Digital Future, you don’t have to wait till it’s been through review and formal publication to read it. I submitted to PeerJ, and at the same time, made it available as a preprint (Taylor 2014).

It’s called “Quantifying the effect of intervertebral cartilage on neutral posture in the necks of sauropod dinosaurs”, and frankly the results are weird. Here’s a taste:

Taylor (2014:figure 3). Effect of adding cartilage to the neutral pose of the neck of Apatosaurus louisae CM 3018. Images of vertebra from Gilmore (1936:plate XXIV). At the bottom, the vertebrae are composed in a horizontal posture. Superimposed, the same vertebrae are shown inclined by the additional extension angles indicated in Table 1. If the slightly sub-horizontal osteological neutral pose of Stevens and Parrish (1999) is correct, then the cartilaginous neutral pose would be correspondingly slightly lower than depicted here, but still much closer to the elevated posture than to horizontal. (Note that the posture shown here would not have been the habitual posture in life: see discussion.)

Taylor (2014:figure 3). Effect of adding cartilage to the neutral pose of the neck of Apatosaurus louisae CM 3018. Images of vertebra from Gilmore (1936:plate XXIV). At the bottom, the vertebrae are composed in a horizontal posture. Superimposed, the same vertebrae are shown inclined by the additional extension angles indicated in Table 1. If the slightly sub-horizontal osteological neutral pose of Stevens and Parrish (1999) is correct, then the cartilaginous neutral pose would be correspondingly slightly lower than depicted here, but still much closer to the elevated posture than to horizontal. (Note that the posture shown here would not have been the habitual posture in life: see discussion.)

A year back, as I was composing a blog-post about our neck-cartilage paper in PLOS ONE (Taylor and Wedel 2013c), I found myself writing down the rather trivial formula for the additional angle of extension at an intervertebral joint once the cartilage is taken into account. In that post, I finished with the promise “I guess that will have to go in a followup now”. Amazingly it’s taken me a year to get that one-pager written and submitted. (Although in the usual way of things, the manuscript ended up being 13 pages long.)

To summarise the main point of the paper: when you insert cartilage of thickness t between two vertebrae whose zygapophyses articulate at height h above the centra, the more anterior vertebra is forced upwards by t/h radians. Our best guess for how much cartilage is between the adjacent vertebrae in an Apatosaurus neck is about 10% of centrum length: the image above shows the effect of inserting that much cartilage at each joint.

And yes, it’s weird. But it’s where the data leads me, so I think it would be dishonest not to publish it.

I’ll be interested to see what the reviewers make of this. You are all of course welcome to leave comments on the preprint itself; but because this is going through conventional peer-review straight away (unlike our Barosaurus preprint), there’s no need to offer the kind of detailed and comprehensive comment that several people did with the previous one. Of course feel free if you wish, but I’m not depending on it.

References

Gilmore Charles W. 1936. Osteology of Apatosaurus, with special reference to specimens in the Carnegie Museum. Memoirs of the Carnegie Museum 11:175–300 and plates XXI–XXXIV.

Stevens, Kent A., and J. Michael Parrish. 1999. Neck posture and feeding habits of two Jurassic sauropod dinosaurs. Science 284(5415):798–800. doi:10.1126/science.284.5415.798

Taylor, Michael P. 2014. Quantifying the effect of intervertebral cartilage on neutral posture in the necks of sauropod dinosaurs. PeerJ PrePrints 2:e588v1 doi:10.7287/peerj.preprints.588v1

Taylor, Michael P., and Mathew J. Wedel. 2013c. The effect of intervertebral cartilage on neutral posture and range of motion in the necks of sauropod dinosaurs. PLOS ONE 8(10):e78214. 17 pages. doi:10.1371/journal.pone.0078214

SO close

August 21, 2014

Bipedal Diplodocus USNM 10865 - modified from Gilmore 1932 pl 6 - v2

I have often argued that given their long hindlimbs, massive tail-bases, and posteriorly-located centers of mass, diplodocids were basically bipeds whose forelimbs happened to reach the ground. I decided to see what that might look like.

Okay, now obviously I know that there are no trackways showing sauropods actually getting around like this. It’s just a thought experiment. But given how close the center of mass of Diplodocus is to the acetabulum, I’ll bet that this pose was achievable in life. If diplodocids had just pushed the CM a few cm farther back, they might have dispensed with forelimbs entirely, or done something different with them, like re-evolved grasping hands.

Image modified from Gilmore (1932: plate 6). Here’s a horizontal-necked bipedal Diplodocus and the original pose:

Bipedal Diplodocus USNM 10865 - modified from Gilmore 1932 pl 6

Diplodocus USNM 10865 - Gilmore 1932 pl 6 - cleaned up

UPDATE the next day: I had forgotten that Niroot had already done a bipedal Apatosaurus, and a much more convincing one than mine. Go see it.

UPDATE the next week: Well, heck. Looks like the primary value of this post was so that people would remind me of all the other places the same idea has already been covered better. As you can see from the comment thread, Mike blogged about this at the WWD site, Scott Hartman drew it, and Heinrich Mallison showed that it was plausible. Sheesh, I suck.

Reference

  • Gilmore, C. W. 1932. On a newly mounted skeleton of Diplodocus in the United States National Museum. Proceedings of the United States National Museum 81, 1-21.

Supersaurus vs Brachiosaurus - BYU 9024 and FMNH P25107

This was inspired by an email Mike sent a couple of days ago:

Remind yourself of the awesomeness of Giraffatitan:
http://svpow.files.wordpress.com/2008/11/mike-by-jango-elbow.jpeg

Now think of this. Its neck is 8.5m long. Knock of one measly meter — for example, by removing one vertebra from the middle of the neck — and you have 7.5 m.

Supersaurus’s neck was probably TWICE that long.

Holy poo.

I replied that I was indeed freaked out, and that it had given me an idea for a post, which you are now reading. I didn’t have a Giraffatitan that was sufficiently distortion-free, so I used my old trusty Brachiosaurus. The vertebra you see there next to Mike and next to the neck of Brachiosaurus is BYU 9024, the longest vertebra that has ever been found from anything, ever.

Regarding the neck length of Supersaurus, and how BYU 9024 came to be referred to Supersaurus, here’s the relevant chunk of my dissertation (Wedel 2007: pp. 208-209):

Supersaurus is without question the longest-necked animal with preserved cervical material. Jim Jensen recovered a single cervical vertebra of Supersaurus from Dry Mesa Quarry in western Colorado. The vertebra, BYU 9024, was originally referred to “Ultrasauros”. Later, both the cervical and the holotype dorsal of “Ultrasauros” were shown to belong to a diplodocid, and they were separately referred to Supersaurus by Jensen (1987) and Curtice et al. (1996), respectively.

BYU 9024 has a centrum length of 1378 mm, and a functional length of 1203 mm (Figure 4-3). At 1400 mm, the longest vertebra of Sauroposeidon is marginally longer in total length [see this post for a visual comparison]. However, that length includes the prezygapophyses, which overhang the condyle, and which are missing from BYU 9024. The centrum length of the largest Sauroposeidon vertebra is about 1250 mm, and the functional length is 1190 mm. BYU 9024 therefore has the largest centrum length and functional length of any vertebra that has ever been discovered for any animal. Furthermore, the Supersaurus vertebra is much larger than the Sauroposeidon vertebrae in diameter, and it is a much more massive element overall.

Neck length estimates for Supersaurus vary depending on the taxon chosen for comparison and the serial position assumed for BYU 9024. The vertebra shares many similarities with Barosaurus that are not found in other diplodocines, including a proportionally long centrum, dual posterior centrodiapophyseal laminae, a low neural spine, and ventrolateral flanges that connect to the parapophyses (and thus might be considered posterior centroparapophyseal laminae, similar to those of Sauroposeidon). The neural spine of BYU 9024 is very low and only very slightly bifurcated at its apex. In these characters, it is most similar to C9 of Barosaurus. However, theproportions of the centrum of BYU 9024 are more similar to those of C14 of Barosaurus, which is the longest vertebra of the neck in AMNH 6341. BYU 9024 is 1.6 times as long as C14 of AMNH 6341 and 1.9 times as long as C9. If it was built like that of Barosaurus, the neck of Supersaurus was at least 13.7 meters (44.8 feet) long, and may have been as long as 16.2 meters (53.2 feet).

Based on new material from Wyoming, Lovelace et al. (2005 [published as Lovelace et al. 2008]) noted potential synapomorphies shared by Supersaurus and Apatosaurus. BYU 9024 does not closely resemble any of the cervical vertebrae of Apatosaurus. Instead of trying to assign its serial position based on morphology, I conservatively assume that it is the longest vertebra in the series if it is from an Apatosaurus-like neck. At 2.7 times longer than C11 of CM 3018, BYU 9024 implies an Apatosaurus-like neck about 13.3 meters
(43.6 feet) long.

Supersaurus vs Diplodocus BYU 9024 and USNM 10865 - Gilmore 1932 pl 6

Bonus comparo: BYU 9024 vs USNM 10865, the mounted Diplodocus longus at the Smithsonian, modified from Gilmore 1932 (plate 6). For this I scaled BYU 9024 against the 1.6-meter femur of this specimen.

If you’d like to gaze upon BYU 9024 without distraction, or put it into a composite of your own, here you go:

Supersaurus cervical BYU 9024

References

 

Check out this beautiful Lego Diplodocus:

10954093715_c4c7fe19ec_k-crop

(Click through for the full image at full size.)

I particularly like the little touch of having of bunch of Lego Victorian gentleman scientists clustered around it, though they’re probably a bit too big for the skeleton.

This is the work of MolochBaal, and all rights are reserved. You can see five more views of this model in his Flickr gallery. I especially admire how he’s managed to get the vertebral transitions pretty smooth, the careful use of separate radius/ulna and tibia/fibula, and the use of a transparent brick in the skull to represent the antorbital fenestra.

The forefeet are wrong — their toes should not be splayed out — but you can’t blame MolochBaal for that, as he was copying the mounted CM 84/94 cast in the Madrid museum.

 

OLYMPUS DIGITAL CAMERA

Now considered a junior synonym of Supersaurus, on very solid grounds.

Incidentally, unlike the neural spines of most non-titanosaurian sauropods, the neural spine of this vertebra is not simply a set of intersecting plates of bone. It is hollow and has a central chamber, presumably pneumatic. Evidence:

OLYMPUS DIGITAL CAMERA

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